Comparison of geographic distribution models of white-tailed deer Odocoileus virginianus...

8/7/2019 Comparison of geographic distribution models of white-tailed deer Odocoileus virginianus (Zimmermann, 1780) sub

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THERYA, Abril, 2010Vol.1(1): 41-68

Abstract

Salvador Mandujano1*, Christian A. Delfn-Alfonso2,3and Sonia Gallina1

The white-tailed deer (Odocoileus virginianus) is the most widely distributed and studiedungulate in the American continent. This species is found throughout Mexico, excepton the peninsula of Baja California and some areas of northern Chihuahua and Sonora.In this study we compared three geographic distribution models (Kellogg 1956; Hall1981; Villarreal 1999) of white-tailed deer subspecies on a national scale, by state andby principal vegetation types. We found that neither the number of subspecies (13 or14 of the 38 recognised subspecies), nor the geographical limits between subspeciescoincided completely between models. Furthermore, for several subspecies, marked

differences in distribution area were found depending on the distribution model used.Using multivariate analyses, we found that the 14 subspecies can be separated into threegroups associated with different vegetation types: the northern subspecies associatedwith shrub land, the Pacic subspecies associated with temperate forest and tropical

dry forest, and the south-eastern subspecies associated with tropical evergreen forest,cloud forest and tropical semi-deciduous forest. We suggest the classication of the 14

subspecies into three ecoregions. The data analyzed here is relevant to the managementand conservation of the white-tailed deer subspecies and/or geographical variations inMexico; it is also important to avoid the translocation of individuals into inappropriateareas with respect to their evolution and adaptation to different ecoregions.

Key words: distribution, ecoregions, management, Mexico, subspecies, vegetationtypes.1 Red de Biologa y Conservacin de Vertebrados, Instituto de Ecologa A. C., Carretera Antigua Coatepec No. 351, ElHaya, Xalapa 91070, Ver., Mxico. E-mail: [email protected], autor corresponsal2 Red de Medio Ambiente y Sustentabilidad, Instituto de Ecologa A. C., Carretera AntiguaCoatepec No. 351, El Haya, Xalapa 91070, Ver., Mxico.3 Facultad de Ciencias Naturales, Universidad Autnoma de Quertaro, Av. de la Ciencia S/N Juriquilla, Delegacin SantaRosa Juregui, Santiago de Quertaro, C. P. 76230, Quertaro, Mxico.

Comparison of geographic distributionmodels of white-tailed deer

Odocoileus virginianus (Zimmermann, 1780)subspecies in Mexico: biological

and management implications


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42 THERYA Vol.1(1): 41-68

WHITE-TAILED DEER SUBSPECIES IN MEXICO

Introduction

El venado cola blanca (Odocoileus virginianus) es el ungulado con mayor reade distribucin en el continente Americano. En Mxico, se le encuentra en todo elterritorio excepto la pennsula de Baja California, algunas reas del norte de Chihuahuay norte de Sonora. En este trabajo comparamos algunos modelos (Kellogg 1956; Hall

1981; Villarreal 1999) de distribucin geogrca de las subespecies de venado colablanca, a nivel del pas, por entidad federativa y por tipos principales de vegetacin. Seencontr que no coinciden totalmente ni en el nmero de subespecies (13 14 de las38 subespecies reconocidas), ni en los lmites geogrcos entre subespecies; adems

de que para algunas subespecies existen claras diferencias en las reas de distribucindependiendo del modelo de distribucin. Con base en anlisis multivariados encontramosque las 14 subespecies pueden separarse en tres grupos asociados a diferentes tiposde vegetacin: las subespecies norteas asociadas al matorral xerlo, las subespecies

del Pacco asociadas al bosque templado y bosques tropical seco, y las subespecies

del sureste asociadas al bosque tropical perennifolio, bosque meslo y bosque

tropical subcaducifolio. Sugerimos clasicar a las subespecies en tres ecoregiones. Lainformacin generada es relevante para el manejo y conservacin de las subespecies y/ovariaciones geogrcas del venado cola blanca en el pas, y evitar translocar individuos

a sitios que no les corresponden de acuerdo a su evolucin y adaptacin a las diferentesecoregiones.

Palabras clave: distribucin, ecoregiones, manejo, Mxico, subespecies, tipos devegetacin.

The white-tailed deer Odocoileus virginianus (Zimmermann 1780) is the most widelydistributed and studied cervid in the American continent (Gallina et al. 2010). It is foundfrom a latitude of 60 N in the south of Canada, through most of the United States, exceptsome regions of the southeast, throughout Central America, and into South America inthe north of Brazil and south of Peru at a latitude of 15 S (Smith 1991; Gallina et al.2010). In Mexico this species is found throughout the country, except on the peninsulaof Baja California and in some areas of northern Chihuahua and Sonora (Leopold 1959).

The high levels of reproductive, behavioural and ecological plasticity observed in thisspecies, are factors that have allowed it to expand its geographic distribution (Baker1984). As a consequence, this browser cervid inhabits an extensive variety of differentplant communities. In Mexico this species is found in temperate pine, oak and r forests,

mixed oak pine forest, shrub land, tropical dry forest, semi-evergreen and evergreenforests, subaquatic vegetation and secondary vegetation (Galindo-Leal and Weber

2005).Thirty eight subspecies of the white-tailed deer have been described, 14 of which are

found in Mexico (Smith 1991). Although the level of subspecies is frequently employed forconservation and management purposes, from a biological point of view its denition is

controversial. Theoretically, subspecies are groups of local populations, within a species,that share a geographical range and common characteristics (genetic and phenotypic),

Resumen


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Salvador Mandujano, Christian A. Delfn-Alfonso and Sonia Gallina

that are adapted to the environmental conditions found in their habitat, and that areseparated from other populations by some kind of geographical or climatic barrier; suchqualities allow for the distinction of one subspecies from another (Frankham et al. 2002).However, the classication of the currently recognised white-tailed deer subspecies is

entirely based on morphological characteristics (e.g., size, pelage colour, size and shapeof male antlers), from just a few museum specimens (Kellogg 1956), and only a limited

studies exist that present detailed or quantitative morphological or genetic data (e. g.,Krausman et al. 1978; Shefeld et al. 1985; Cronin 1991a, 1991b; Mathews and Porter1993; Ellsworth et al.1994; Anderson et al. 2002; Van Den Bussche et al. 2002; DeYounget al. 2002, 2003). For Mexico, very little information about morphometric or geneticvariability in white-tailed deer is available. Studies exist for four north-eastern subspecies(Logan-Lopez et al. 2006, 2007), ve subspecies in the country (Caldern-Lobato 2009),

and three subspecies in the state of Michoacan (Chassin, personal communication). Forthese reasons, there is a clear need for further research in this topic.

The range of individual variation found in white-tailed deer, especially in size,antler details, and pelage colour, is remarkable and has been used for their subdivision

into subspecies (Kellogg 1956). However, one form can pass gradually into another,especially where there are no abrupt changes in physiography. Specimens from certainregions might, with equal propriety, be referred to as either of the contiguous subspecies.But where abrupt changes in physiography do occur, such as the change from plainsto high mountains, the deer tend to respond along rather sharp lines of differentiation.Generally, among the subspecies of white-tailed deer, the larger forms are found in thenorth and the smaller forms in the south. For example, the maximum size is reachedin those races extending westward from the Atlantic coast across southern Canadaand northern United States. The minimum size is found in O. v. rothschildi,inhabitingCoiba Island off the Pacic coast of Panama. In Mexico, the largest subspecies is O. v.

texanus,while the smallest is O. v. acapulcensis. Variation in antler-formation seemsto correlate mainly with various physical factors, among which are size, maturity, andgeneral physical condition (Villarreal 1999). The larger northern subspecies carry largerantlers, with more numerous tines, than do the smaller southern forms. For example,in O. v. acapulcensis the antlers, even in fully mature bucks, may be reduced to singlespikes; in contrast, in O. v. texanus subsidiary tines are more prevalent. Individual andsubspecic variations in colour are so great that they are perplexing to explain, and

gradations are so numerous that they are difcult to distinguish. The pelages usually are

quiet different among seasons. For example, in the northern subspecies the winter andsummer colour differ, while in Mexico there are variations between wet and dry seasons.In general, summer or wet season colour being predominantly brownish or greyish, whilewinter or dry season is tawny. The most vivid coloration, ranging from tawny to richorange-cinnamon, is shown by some of the subspecies inhabiting the tropical lowlandsin Mexico and Central America. The colours of individuals from the high mountainsof these regions usually are dull and dark. A further factor confusing the situation isthe widespread translocation of various subspecies of white-tails into geographic rangesproperly belonging to others.

The white-tailed deer is a highly valued species in Mexico for many reasons,including: protein consumption, commerce, the elaboration of artisan crafts, and as an


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important component in the rituals of indigenous cultures (Mandujano and Rico-Gray1991; Greenberg 1992; Gonzalez-Perez and Briones 2000; Naranjo et al. 2004). Todaythe hunting of this species continues to be important, both for subsistence and trophyhunting. The white-tailed deer is the most important game species in Mexico and itscynegetic exploitation has increased notably in Wildlife Conservation, Managementand Sustainable Utilization Units (in Spanish: Unidades de Manejo y Aprovechamiento

para la Conservacin de la Vida Silvestre, or UMAs, Montiel et al. 1999; Gonzalez-Marin et al. 2003; Segovia and Hernandez 2003; Villarreal-Espino 2006), providing animportant source of income, particularly in the north of the country (Villarreal 1999). Inconsequence, UMA management has led to a greater national demand for the hunting ofwhite-tailed deer and an urgent need for reliable biological and ecological data in orderto sustainably manage the different populations and subspecies. In this sense, NaturalProtected Areas (in Spanish ANPs) have proven to be important areas for such studies(Gallina et al. 2007).

From a cynegetic management point of view, in Mexico only ve of the 14 white-

tailed deer subspecies enter into the current international trophy record books, such

as those organised by the Boone and Crockett Club and Safari Club International. Forthis reason, these are the subspecies that are given the most attention and protectionby ranchers and land owners. These subspecies are: O. v. texanus,O. v. couesi, O. v.carminis, O. v. miquihuanensis and, in recent years, O. v. mexicanus (Villarreal-Espino2002). The remaining subspecies have not been considered as recognised trophies bynational or overseas sport hunters, owing both to their smaller sized antlers and their lackof an individual category in the trophy record books. Notwithstanding, the situation isbeginning to change and a greater regional importance is being placed on the huntingof each individual subspecies (Villarreal 2009). As a result, one worrying aspect ofwhite-tailed deer management is the translocation of subspecies to different parts of the

country other than those areas where they are historically found. Given this problem,the Wildlife Department of SEMARNAT (the Mexican ofce for the Environment andNatural Resources) requires that any animals that are translocated are of the samesubspecies as those found in the local area. This legislation forms part of Article 81of the General Wildlife Law of Mexico (Diario Ocial de la Federacin 2006). Thisis particularly important in large UMAs, where animals are often released with littlecontrol. For this reason there is an urgent need for a database of geographic informationthat allows both the authorities and farm owners to gain a better understanding of thesubspecies they manage, and to maintain the genetic variability of the species and, inturn, its conservation prospects. See Appendix 1 to detail description of each subspeciesin Mexico.

Based on information from the Smithsonian Institution, Kellogg (1956) publishedan interesting map, now little known, in Taylors (1956) book, which has been considereda classic for decades. Interestingly, the author only recognised 13 white-tailed deersubspecies in Mexico and presented geographical limits between subspecies which werediscontinuous. That is, he left empty spaces in which it is difcult to dene which

subspecies are distributed (Fig. 1a). The distribution map by Hall (1981) presents thecontinuous distribution of 14 subspecies, based on records from limits of the distributionof each subspecies (Fig. 1b). This map appears to be the basis of others, such as that


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Figure 1. Distributionmodels of the white-tailed deer subspeciesin Mexico, proposedby Kellogg (1956), Hall(1981) and Villarreal(1999). ? Indicates theregions with doubt asto which subspeciesare present. In the threemodels, each subspeciesis presented by the samecolour.


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Methods

published by Baker (1984) which, in turn, served as the basis of the map presentedby Smith (1991) in a monograph of the species, a mandatory reference. For Mexico,Villarreal (1999) published a distribution map of the subspecies which contains someimportant differences with respect to the other maps previously mentioned (Fig. 1c).This book is frequently consulted by deer managers in Mexico, and the author basedthe map on that proposed by Baker (1984), but introduced signicant changes based

on eld experience with the species. How comparable are these distribution models?Could differences be expected in the number of subspecies and distribution surfacein each federal entity depending of the distribution model used? Considering not thetaxonomic characteristics, could the 14 subspecies be classied different according

with the ecological conditions where they inhabit? Which are the conservation andmanagement implications of this distribution models comparison? The objective of thisstudy was to compare the biogeographic distribution maps for the subspecies of white-tailed deer in Mexico (Kellogg 1956; Hall 1981; Villarreal 1999), in order to: 1) Estimatethe distribution areas of the subspecies on a national scale, by state and by vegetationtype. 2) Identify related groups of subspecies in function with the principal vegetation

types found in the country in order to dene possible ecoregions, and 3) Analyse theimplications of our results for the biology and management of the 14 subspecies found inMexico. Since a model is a graphical, mathematical, physical, or verbal representationor simplied version of a concept, phenomenon, relationship, structure, system, or any

aspect of the real world, and a species distribution map is a visual representation in whicha biological taxon is spatially arranged in an area, in this paper we used distributionmodel as a synonym for distribution map.

Sources of information used.- Information relating to the distribution of the subspecies

of white-tailed deer in Mexico was gathered by consulting the maps of Kellogg (1956),Hall (1981), and Villarreal (1999). Information additional was consulted in Goldmanand Kellogg (1940), Miller and Kellogg (1955), Hall and Kelson (1959), Taylor (1956),Rue (1979), Hall (1981), Smith (1991), Villarreal (1999) and Heffelnger (2006). We

also obtained the bibliographical databases published in Mandujano (2004) and Gallinaet al. (2008, 2009). Biogeographic models of physiographic regions and vegetation inMexico were taken from Rzedowski and Reina-Trujillo (1990), Arriaga et al. (1997),Palacio et al. (2000) and Morrone (2005). Further, several digital maps of the 32 federalentities of Mexico (31 states and one Federal District; hereinafter states) were obtainedfrom several sources, including: State Political Divisions of INEGI (National Institute ofStatistics and Geography), Biogeographic Regions (Arriaga et al. 1997), National ForestInventory (Palacio et al. 2000), and Floristic Divisions (Rzedowski and Reyna-Trujillo1990). All of the maps on which this study was based were obtained from the databasesif INEGI (http://mapserver.inegi.org.mx) and CONABIO (National Commision for theUnderstanding and Use of Biodiversity, http://www.conabio.gob.mx/informacion/gis/).

Analysis of geographic and statistical data.- The three distribution maps used weredigitalised using Arc View ver. 3.2 (ESRI 1999) and homogenised into a plane coordinatesystem in order to simplify the area calculations into. The metadata used were from a map


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generated by CONABIO (http://www.conabio.gob.mx/informacion/). The resulting mapswere geo-referenced to allow for simple, comparative referencing. Each of the distributionmodels was broken down into section by state and vegetation type, as mentioned earlier.Later, the areas of the different sections were calculated for each distribution model: the

total area occupied by each subspecies, the area occupied by each subspecies in eachstate, and the area occupied by each subspecies in each vegetation type.

With the data we generated with regard to the area occupied by each subspecieswithin each vegetation type in each distribution model, we obtained percentages for eachsubspecies. With the percentage data for each vegetation type, we carried out ClusterAnalyses (with the algorithm Euclidean Paired Group Similarity Index and PrincipalComponent Analyses (PCA)), using correlation matrices and Euclidean Similarity Indexfor each distribution model using, in both cases, the statistics program PAST (Hammer etal. 2001). The results of both multivariate analyses allow related groups of subspecies,sharing similar vegetation types, to be dened. We dened these groups as possible

ecoregions.Based on the studies by Kellogg (1956) and Villarreal (2009), in Appendix 1. We

show a description of the subspecies found in Mexico. However, it is important tonote that the data presented are based on measurements taken from one, or just a fewindividuals. In the case ofO. v. toltecus no data are available. We analysed the datafrom Appendix 1 using PCA with a correlation matrix, in order to know the relationshipsof the 13 Mexican subspecies (excluding O. v. toltecus, for which data was unavailable)in function of their size.

Geographic distribution of subspecies.-The total distribution area for any given subspeciesdiffered markedly, depending on the distribution model used (Table 1). The subspecies

that differed in their distribution area, depending on the model were: O. v. carminis,O. v. oaxacensis, O. v. miquihuanensis, O. v. nelsoni, O. v. toltecus, O. v. veraecrucisand O. v. yucatanensis. If we do not consider the model used by Kellogg (1956), thatunderestimates distribution area, the subspecies with the greatest distribution area wasO. v. couesi, which inhabits >500,000 km2, followed by O. v. miquihuanensis (~200,000km2), O. v. sinaloae, O. v. mexicanus, O. v. texanus, and O. v. thomasi (~172,000 to106,000 km2). Other subspecies that varied between 60,000 and 106,000 km2 were:O v. carminis, O. v. veraecrucis, O. v. yucatanensis and O. v. acapulcensis. Finally, theremaining subspecies, O. v truei, O. v toltecus, O. v. oaxacensis, O. v. nelsoni, and werefound to have the most restricted distribution areas (< 62,000 km2).

Distribution by federal entity.- The number of subspecies that could apparently be foundin each state (federal entity) varied depending on the distribution model used, primarilyvarying with the use of the Kellogg (1956) model, which could be considered over-conservative (Table 2). The state with the greatest number of subspecies was Oaxaca,with 5 or 6, followed by Durango, Guanajuato, Guerrero, Jalisco and Veracruz with 3or 4. Those states that contained only one subspecies were: Distrito Federal, Estado deMxico, Morelos, Tabasco, Tlaxcala and Yucatn. Of the 30 states considered, only in13 did the number of subspecies found coincide among the three models. In the case

Results


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of the state of Aguascalientes, in Kelloggs (1956) model no subspecies are reported tobe found because is considered problematic to dene the geographic limits between

subspecies (Fig. 1).The subspecies that inhabited the greatest number of states was O. v. mexicanus (13 of

30, including Distrito Federal), followed by O. v. couesi, O. v. miquihuanensis and O.

v. sinaloae. The only subspecies limited to just one state was O. v. nelsoni; while O. v.oaxacensis and O. v. trueiwere limited to two states (Fig. 1). Although we do not presentthe data, given the differences in the distribution of subspecies between models, the areainhabited by each subspecies in each state varied considerably.

Distribution by vegetation type.- It is important to note that of the 14 subspecies foundin Mexico, only two (O. v. texanus and O. v. carminis) did not inhabit tropical forests;while all of the other 12 subspecies included at least some tropical forest within theirdistribution areas (Fig. 2 and Table 3). The subspecies O. v. miquihuanensis and O. v.couesi, considered to be principally found in shrub land and temperate forest respectively,were also found to marginally inhabit dry tropical forests. However, those subspecieswith a greater part of their total distribution area occupied by deciduous or tropical dryforests were: O. v. sinaloae, O. v. mexicanus, O. v. acapulcensis and O. v. yucatanensis.In contrast, the subspecies O. v. veraecrucis, O. v. toltecus, O. v. thomasi, O. v. nelsoniand O. v. truei, inhabited tropical evergreen forest, tropical semi-deciduous forest andcloud forest.

SubspeciesDistribution Model

Kellogg+ Hall Villarreal

O. v. acapulcensis 66,014 69,248 64,077

O. v. carminis 26,315 94,031 102,390

O. v. couesi 451,518 548,802 528,928

O. v. mexicanus 121,544 171,574 143,697

O. v. miquihuanensis 94,250 175,221 223,362

O. v. nelsoni 38,142 40,500 27,750

O. v. oaxacensis 44,751 7,820 30,223

O. v. sinaloae 86,047 135,536 168,225O. v. texanus 147,558 167,518 141,026

O. v. thomasi 76,939 105,737 117,363

O. v. toltecus 24,654 62,186 32,455

O. v. truei - 26,378 46,189

O. v. veraecrucis 37,296 76,257 95,505

O. v. yucatanensis 127,965 84,612 71,753

Table 1. Surface (km2)

occupied by each ofthe white-tailed deersubspecies, dependingon the distribution modelused. + According withthe delimitation of thegeographic distributionof each subspecies, thismodel sub-estimatedthe distribution surfacefor some subspecies; seetext.


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Groups of subspecies by vegetation type.- Both PCA (Fig. 3) and Cluster Analysis (Fig.4) gave similar results for all three distribution models. In general, PCA revealed a clearorder of the subspecies O. v. texanus, O. v. miquihuanensis, O. v. carminis and O. v.couesi in function with the presence of shrub land in the north of Mexico; althoughO. v. couesi is more clearly associated with temperate forests. The subspecies O. v.mexicanus, O. v. sinaloae, O. v. oaxacensis and O. v. acapulcensis formed a groupassociated with temperate pine-oak forest and tropical dry forest, mostly in the Pacic

region and the centre of the country. The subspecies O. v. veraecrucis, O. v. thomasi, O.

Table 2. Distribution ofthe white-tailed deersubspecies in eachstate or federal entity inMexico. + In these states,the number of subspeciesis the same in the threemodels used

State

Distribution model

Kellogg Hall Villarreal

Aguascalientes cou, miq cou, miq

Campeche tho, yuc tho, tru, yuc tho, tru, yuc

Chiapas+ nel, tho nel, tho nel, tho

Chihuahua+ car, cou, tex car, cou, tex car, cou, tex

Coahuila car, cou, miq, tex car, miq, tex car, miq, tex

Colima aca, sin aca, sin sin

Distrito Federal+ mex mex mex

Durango cou, sin car, cou, miq, sin car, cou, miq, sin

Guanajuato mex cou, mex, miq, sin cou, mex, miq, sin

Guerrero aca, mex, oax aca, mex, sin aca, mex, oax, sin

Hidalgo mex mex, ver mex, ver

Jalisco aca, cou, mex, sin aca, cou, miq, sin cou, miq, sin

Mxico+ mex mex mex

Michoacn+ aca, mex, sin aca, mex, sin aca, mex, sin

Morelos+ mex mex mex

Nayarit sin cou, sin cou, sin

Nuevo Len+ miq, tex miq, tex miq, tex

Oaxaca aca, oax, tho, tol aca, mex, oax, tho,tol aca, mex, oax, tho, tol, ver

Puebla mex, oax, tol, ver mex, tol, ver mex, tol, ver

Quertaro mex mex, ver mex, miq, ver

Quintana Roo yuc tru, yuc tru, yuc

San Luis Potos mex, miq mex, miq, ver mex, miq, ver

Sinaloa+ cou, sin cou, sin cou, sin

Sonora cou cou cou, sin

Tabasco+ tho tho tho

Tamaulipas+ miq, tex, ver miq, tex, ver miq, tex, ver

Tlaxcala+ mex mex mex

Veracruz tho, tol, ver mex, tho, tol, ver mex, tho, tol, ver

Yucatn+ yuc yuc yuc

Zacatecas+ cou, miq cou, miq cou, miq


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Figure 2. Distributionmodels of the white-tailed deer subspeciesin Mexico within theprincipal vegetationtypes proposed byRzedowski and Reyna-Trujillo (1990).

a).-Kellogg model

b).-Hall model

c).-Villarreal model


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Subspecies Vegetation types

Distribution model


O. v.acapulcensis Temperate forest

Thorn forestCloud forestTropical dry forestTropical semi-deciduous forestTropical rainforestGrasslandShrub landAquatic vegetation

26,038

9691,22126,87410,002

-207

--

26,592

2,4081,22528,9868,691

-312

--

29,751156

1,22123,6928,894

-384

--

O. v. carminis Temperate forestThorn forestCloud forestTropical dry forestTropical semi-deciduous forestTropical rainforestGrasslandShrub landAquatic vegetation

783-----

1,03724,444

-

1,429-----

4,10788,487

-

1,585-----

7,02193,359

-

O. v. couesi Temperate forestThorn forestCloud forestTropical dry forestTropical semi-deciduous forestTropical rainforiestGrasslandShrub landAquatic vegetation

146,33957,047-

31,342--

100,917115,426

-

173,89156,989-

57,725--

110,364149,177

-

173,91242,842-

46,453--

120,832143,746

-

O. v.mexicanus

Temperate forestThorn forestCloud forestTropical dry forestTropical semi-deciduous forestTropical rainforestGrasslandShrub landAquatic vegetation

41,5257,461

61430,406

-658

9,60129,1392,140

66,4156,5011,970

55,707811

4,1834,833

28,9552,198

46,1904,5921,738

53,063-

5,6803,995

26,1952,245

O. v.miquihuanensis


14,841-----

1,78377,626

-

18,2591,608

1422,038

350-

12,839139,986

-

20,598375136

1,014188

-7,881

193,169-

O. v. nelsoni Temperate forestThorn forestCloud forestTropical dry forestTropical semi-deciduous forestTropical rainforestGrasslandShrub landAquatic vegetation

15,091-

1,2799,850

-11,922

---

15,814-

94910,749

2412,747

---

8,958-

1926,950

-11,911

---

O. v.oaxacensis


20,823-

1,02917,526

-221199

4,953-

5,232-

1052,483

-----

20,31316

1969,495

--

37166

-

Table 3. Distributionpotential (km2) ofthe white-tailed deersubspeceis in each ofthe vegetation types byRzedowski and Reyna-Trujillo (1990) andmodied by CONABIO.


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Subspecies Vegetation types

Distribution model


O. v. sinaloae Temperate forestThorn forestCloud forest

Tropical dry forestTropical semi-deciduous forestTropical rainforestGrasslandShrub landAquatic vegetation

27,6479,584

668

41,4434,984---

1,182

39,76512,455

683

70,8616,221-

2,854-

1,996

43,08229,633

684

85,2086,862-

784-

2,279

O. v. texanus Temperate forestThorn forestCloud forestTropical dry forestTropical semi-deciduous forestTropical rainforestGrasslandShrub landAquatic vegetation

9037,239

----

2,980135,504

-

1,1118,820

-237

--

2,787153,911

-

4398,426

----

12,960117,857

-

O. v. thomasi Temperate forestThorn forestCloud forest

Tropical dry forestTropical semi-deciduous forestTropical rainforestGrasslandShrub landAquatic vegetation

5,2401,5442,130

5,3534,725

41,7392,441

-11,814

6,7131,9513,675

5,6874,998

65,7252,844

-11,791

13,5012,1603,724

9,7668,092

65,1342,932

-12,183

O. v. toltecusTemperate forestThorn forestCloud forestTropical dry forestTropical semi-deciduous forestTropical rainforestGrasslandShrub landAquatic vegetation

4,419-

3,4411,000

-13,667

-1,919

208

13,601-

7,5567,789

-26,767

-6,270

202

5,161826

5,5514,136

-16,768

-12

-

O. v. truei Temperate forest

Thorn forestCloud forestTropical dry forestTropical semi-deciduous forestTropical rainforestGrasslandShrub landAquatic vegetation

-

--------

-

2,222-

430-

19,924--

1,876

-

49,482---

39,729--

881

O. v.veraecrucis


8006,8351,6197,248

-17,302

6571,616

816

3,75117,2681,524

13,140123

28,339674

9,5971,123

9,10918,9344,390

16,869284

31,293970

12,9531,048

O. v.yucatanensis


-4,533

-16,23033,50967,228

50-

3,838

-859

-16,01833,10431,709

--

1,800

-762

-18,24031,73321,401

--

1,594

continuacin Tabla 3...


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v. toltecus, O. v. trueiand O. v. yucatanensis are associated with tropical rain forest, semi-deciduous forest, cloud forest, thorn forest and aquatic and sub-aquatic vegetation in thesoutheast. Finally, the subspecies O. v. nelsoniwas an intermediate subspecies betweenthe temperate and dry forest group and the tropical rainforest and semi-deciduous group.With slight variations, this tendency was observed in all three models (Fig. 3).

Cluster Analysis revealed similar groups of subspecies, with slight differences. The

main difference being that O. v. couesi was associated with the temperate/dry forestgroup, depending on the distribution model used (Fig. 4). Consequently, both PCA andCluster Analysis dened three distinct groups of subspecies, clearly associated with

vegetation types. This classication into three groups can be considered as a proposal for

distinct ecoregions, or possible ecotypes for white-tailed deer in Mexico: the ecoregionthat includes the shrub land of the northeast; that which includes the temperate forestsand tropical dry forests of the Pacic and central country; and that which includes the

tropical rain forests, semi-deciduous forests and cloud forests of the Gulf and southeastof the country.

Size of subspecies.- Analysis by PCA demonstrated that 13 of the 14 subspecies foundin Mexico (excluding O. v. toltecus, for which data was unavailable) can be organisedby size, based on total length, chest height and antler measurement (Fig. 5). The largestsubspecies was O. v. texanus, followed by O. v. carminis, O. v. miquihuanensis, O.v. veraecrucis, O. v. mexicanus and O. v. couesi; while the smallest subspecies wereO. v. sinaloae, O. v. thomasi, O. v. yucatanensis, O. v. truei, O. v. oaxacensis, O. v.

acapulcensis and O. v. nelsoni.

Biological implications.- The maps, or distribution models, proposed by Kellogg (1956),

more than 50 years ago, Hall (1981), which was widely accepted, and Villarreal (1999),which is the most commonly used map in Mexico, are not exempt from errors in theirdenitions of the biogeographical limits between subspecies. To reduce this, Kellogg

(1956) presented distribution areas in which the geographical limits between subspecieswere discontinuous. In contrast, Hall (1981) extended the geographic distribution ofeach subspecies and proposed clearly dened limits. However, these limits were dened

on the basis of the authors own criteria and not necessarily on the basis of quantitativestudies. Further, this author recognised a fourteenth subspecies, O. v. truei(= nemoralis),located in the southeast of the country. Another important difference between themodels is that Hall (1981) signicantly increased the distribution area ofO. v. carminis,which Kellogg (1956) reported as found only in a highly restricted area of the Sierra delCarmen, Coahuila. Morphometric studies ofO. v. carminis by Krausman et al. (1978)conrmed the difference with other subspecies; but its geographical limits its unknown.

By increasing the distribution area, Hall (1981) introduced this subspecies into areas ofshrub land, where O. v. texanus and O. v. miquihuanensis were also found. Anotherdifference between the models is that, while Kellogg (1956) presents a wide-rangingdistribution area for O. v. oaxacensis, Hall (1981) reduces this distribution to a very smallarea of the Valles Centrales of Oaxaca. This means that distinguishing O. v. oaxacensisfrom O. v. toltecus can be complicated and their areas of sympatry can be hard to dene.

Discussion


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54 THERYA Vol.1(1): 41-68


Figure 3. Ordination ofthe subspecies of white-tailed deer in functionwith vegetation typefor each of the threedistribution models. Thevariation explained bythe rst two principalcomponents were

53.4%, 53.2% and54.04% for the Kellogg,Hall and Villarrealmodels, respectively.Abbreviations: shrubland (SL), grassland (G),temperate mixed forest(TMF), tropical dry forest(TDF), tropical humidforest (THF), tropicalsemi-deciduous forest(TSF), cloud forest (CF),thorn forest (TF), semi-aquatic vegetation (AV).

Kellogg model

Hall model

Villarreal model


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Figure 4. Classicationof the subspecies ofwhite-tailed deer infunction with vegetationtype for each of the threedistribution models,following the algorithmEuclidean Paired GroupSimilarity Index. Thecoefcients of correlationwere 0.93, 0.91 and0.87 for the Kellogg, Halland Villarreal models,respectively.

Hall model

Kellogg model

Villarreal model


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56 THERYA Vol.1(1): 41-68


For example, Ortiz-Martnez et al. (2005) and Briones and Garca (2005) identify O. v.oaxacensis for the north of the state of Oaxaca. The map proposed by Villarreal (1999)identies 14 subspecies, but modies their distribution limits. For example, in the case

ofO. v. oaxacensis important changes are made, while the distribution of O. v. sinaloae isextended to the south of Sonora and the distribution limits are modied for O. v. texanus,

O. v. miquihuanensis and O. v. carminis (Fig. 1). These differences among distribution

models were evident when we estimate the area of distribution at national and federalstate level (Table 2). In some cases, even the number of subspecies by federal entity wasdifferent depending of the model used (Table 1). Also, these differences are evident inthe vegetation type occupied by each subspecies (Table 3).

From an ecological perspective, our results suggests that the 14 subspeciescan be grouped into three ecoregions: north-eastern shrub land (O. v. carminis, O. v.miquihuanensis and O. v. texanus), Pacic and central temperate and tropical dry forests(, O. v. acapulcensis, O. v. couesi, O. v. mexicanus, O. v. oaxacensis and O. v. sinaloae,),and Gulf-southeast tropical humid, sub-deciduous and cloud forests (O. v. nelsoni, O. v.thomasi, O. v. toltecus, O. v. truei, O. v. veraecrucis and O. v. yucatanensis). An ecoregion

is dened as a relatively large area of land or water that contains a geographicallydistinct assemblage of natural communities and environmental conditions (WWF 1999).These communities share a large majority of their species, dynamics, and environmentalconditions, and function together effectively as a conservation unit at global andcontinental scales (Dinerstein et al. 1995). Several standard methods of classifyingecoregions have been developed, with climate, altitude, and predominant vegetationbeing important criteria (Olson et al. 2001). Because ecoregions are dened by theirshared biotic and abiotic characteristics, they represent practical units on which to baseconservation planning. For example, a map of Mexican ecoregions was presented byDinerstain et al. (1995).

Figure 5. Ordinationof the white-taileddeer subspecies inMexico in functionwith the morphologicalcharacteristics describedin Table 1. The variationexplained by the rst twoprincipal componentswas 79.9%. The rstcomponent (60%)characterises, in the rightside of the graph, thosesubspecies of greatersize expressed as thetotal length (total), height

of shoulder (height) andantler size (antler).


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Figure 6. Classication ofthe subspecies of white-tailed deer in Mexicoaccording to huntingregions by Villarreal(2009) and ecoregionsproposed by Mandujanoet al. (this paper).

The use of ecoregions for deer classication and management has also been suggested

for mule deer (Heffelnger et al. 2006). The 11 subspecies of mule deer and black-taileddeer are distributed throughout western United States and the northern Mexican states.With this wide latitudinal and geographic range, mule deer occupy a great diversity ofclimatic regimes and vegetation associations, resulting in an incredibly diverse set ofbehavioural and ecological adaptations that have allowed this species to succeed. Within

the geographic distribution of mule deer, however, areas can be grouped together intoecoregions, within which deer populations share certain similarities. With regard to theissues and challenges that deer managers face, deVos et al. (2003) has designated sevenseparate ecoregions: California Woodland Chaparral, Colorado Plateau Shrubland andForest, Coastal Rain Forest, Great Plains, Intermountain West, Northern Boreal Forest,and Southwest Deserts. The diversity among the ecoregions presents different challengesto deer managers and guidelines for managing habitat must address these differences(Heffelnger et al. 2003).

For the distribution of white-tailed deer subspecies in the United States, Deckman(2003) classied the 16 subspecies into six regions: Western Region I (O. v. couesi, O. v.

leucurus, and O. v. ochrourus). North Central Region II (O. v. dacotensis). Central PlainsRegion III (O. v. macrourus and O. v. texanus). Gulf Coast Region IV (O. v. clavium, O.v. mcilhennyi, O. v. osceola and O. v. seminolus). Atlantic Islands and SoutheasternRegion V (O. v. hiltonensis, O. v. nigribarbis, O. v. taurinsulae, O. v. venatorius and O.v. virginianus), and Northeastern Region VI (O. v. borealis). However, this author alsorecognised that white-tailed deer subspecies could overlap in many areas and that thegenetic pool in these areas is likely to be mixed.

For Mexico, the Comisin de Flora y Fauna Silvestres de Nuevo Len (CEFFSNL)

recently proposed to Safari Club International the inclusion of nine hunting regions,with the objective of recognising all of the 14 different subspecies of white-tailed deer inMexico on an international scale as different sport hunting trophies (Villarreal 2009). Thisproposal comes from the fact that the Safari Club International and Boone and Crockett


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58 THERYA Vol.1(1): 41-68


Club only recognise ve subspecies. The hunting regions were dened on the basis of

antler size, in such as way that males of the different subspecies are competitive within anygiven geographic region with particular ecological characteristics. The hunting regionsproposed by Villarreal (2009) are presented in gure 6, in which we also compare the

classication of the ecoregions proposed in this article. This gure represents a synthesis

of both hunting and ecological view of the 14 subspecies in Mexico, and we propose it

as a framework for management and conservation actions.The taxonomic rank of subspecies has been the subject of long-running controversy (Mayr1982). Traditionally, subspecies have been recognised on the basis of discontinuitiesin the geographical distribution of phenotypic traits. It has long been established thatpopulations on islands encounter a physical impediment to gene-ow between local

populations, and it is therefore expected that such populations may diverge in isolation(Mayr and Ashlock 1991). In contrast, continental subspecies will often have geographicalranges that directly adjoin, or even overlap, those of conspecic subspecies, and thus

any phenotypic adaptation to local environments will need to take place in the faceof gene ow (e. g., ecotype). Throughout the geographic range of white-tailed deer in

Mexico, we see a lot of variation in body size, pelage colour, antler shape, and otherattributes (Kellogg 1956, Villarreal 2009). For example, subspecies in the southernlatitudes are generally smaller in body size than those in the north, and those inhabitingopen habitats appear lighter in colour than those in heavily forested habitats. As was thecase with mule deer O. hemionus (Heffelnger 2006), the geographic range of severalwhite-tailed deer subspecies was drawn somewhat arbitrarily. In fact, the purporteddifferences between subspecies were often based on subjective opinions regardingcharacteristics or measurements of only one or a few specimens. Fortunately, recentadvances in DNA analysis techniques now allow researchers to evaluate genetic datain ways that provide managers with meaningful ecological management units on an

ecoregion scale. Therefore, given the lack of quantitative morphometric and genetic datafor the subspecies, it is impossible to conrm the taxonomic validity of the subspeciespresent in the country. Further, the absence of phylogeographic studies means that theredenition of the geographic limits between subspecies is not possible. Indeed, when

faced with a species with such capacity for adaptation to different environments, it ishard to dene where the distribution of one subspecies ends and another begins (i.e.,

zones of sympatry) as, basically, some kind of geographic (mountains, rivers, drasticchanges in vegetation type) or climatic barrier ought to exist in order to clearly dene

subspecies. However, in some cases we may expect to nd a gradient of variations, or

cline, from one subspecies to the next and, therefore, the distribution limits betweenthem become arbitrary.

Management implications.- Given the lack of data supporting the validity of thebiogeographic limits of the subspecies, there should be strict control over the deliberate,or even accidental, movement of subspecies to localities where they have not beenhistorically reported. One important problem is posed by those regions where morethan one subspecies converge and the criteria to dene the geographic limits between

the subspecies are arbitrary. The translocation of animal species has resulted in severeconsequences for conservation, such as interspecic competition and the introduction of


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Acknowledgements

diseases and parasites (Waldrup et al. 1990; Kock et al. 2007). For example, Galindo andWeber (1994) reported incidences of dystocia (difculty in parturition), probably caused

by the translocation of some individuals of other subspecies, such as O. v. texanus, intothe distribution range of O. v. couesi, in La Michila, Durango. Unfortunately, someprogrammes translocate individuals, principally of the subspecies O. v. texanus, to otherparts of the country, with the idea of improving the breed, mainly the size of their

antlers. In such a case, Storfer (1999) suggested that knowledge of gene ow rates andunderstanding ecological differences among populations is necessary before embarkingon a program to articially enhance gene ow. This situation could change in the

medium term. For example, recently, the CEFFSNL stated that although they had always

supported and developed cynegetic activities for the benet of the economy, they would

not agree to support the transfer of individuals of the subspecies O. v. texanus from thenorth of Mexico (Coahuila, Nuevo Len and Tamaulipas) to other parts of the country,

even under regulated management schemes such as in UMAs. This is to avoid crossbreading between subspecies that are adapted to differing ecological conditions and thatform part of the biological heritage of the country (Villarreal 2009). Our classication

of the 14 subspecies into three possible ecoregions suggests the more critical actionto translocate deer among ecoregions than inside of the same ecoregion. Ecoregionsclassication implicates a local adaptation to similar ecological conditions.

From a conservation perspective, Phillimore and Owens (2006) suggest thatsubspecies may be of considerable conservation value, as proxies for the sub-structurefound within species. They suggest that the conservation value of subspecies is likely tobe greatest in situations where molecular data is absent, a scenario that is encountered inthe white-tailed deer in Mexico. However, there is an urgent need to integrate biographicmodels and molecular studies (e. g., Moodley and Bruford 2007) as a framework forthe conservation of white-tailed deer at the national and continental scale. Thus, it is

imperative to obtain data on the geographical variation of white-tailed deer throughoutthe country. The studies must investigate both morphological and genetic variation. Thisevaluation will provide not only a basis for conservation efforts (Gallina and Mandujano2009), but also help solidify the range of different subspecies into and among ecoregions.Clear delineation of the boundary among subspecies is an issue that the governmentalofces (SEMARNAT, DGVS, CONANP, CONABIO) and international agencies such as

Safari Club International must address in order to maintain the integrity of the differentgeographical haplotypes and for trophy record books.

We are grateful to the editor T. lvarez for the invitation to contribute in this inauguraledition of the journal Theyra. T. Prez kindly helped with the organisation of the tables.To J, A. Dunn for check the nal version of the manuscript in English. This study include

the Red de Biologa y Conservacin de Vertebrados and Red de Medio Ambiente ySustentabilidad of the Instituto de Ecologa, A. C.


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Atlas Nacional de Mxico. Vol. II. Instituto de Geografa, Universidad NacionalAutnoma Mxico. Mxico.

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histocompatibility locus within and among populations of white-tailed deer(Odocoileus virginianus). Journal of Mammalogy 83:31-39.

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ViLLArreAL-espino, o. 2006. El Venado Cola Blanca en la Mixteca Poblana: Conceptos yMtodos para su Conservacin y Manejo. Fundacin Produce Puebla A. C., Puebla.A. 2006. El Venado Cola Blanca en la Mixteca Poblana: Conceptos y Mtodos parasu Conservacin y Manejo. Fundacin Produce Puebla A. C., Puebla.

wALdrup, K., A. KocAn, r. BArKer, And G. wAGer.1990. Transmission ofBabesia odocoileiin white-tailed deer by Ixodes scapularis. Journal of Wildlife Diseases 26:390-

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June 19, 2002).

Sometido: 3 octubre 2009

Revisado: 14 enero 2010

Aceptado: 18 febrero 2010Editor asociado Jan Schipper


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66 THERYA Vol.1(1): 41-68


Su

bspec

ies

Type

loca

lity

Measu

remen

ts(mm

)

An

tler

Minimum

score

for

trop

hy

hun

ting

(a=a

typ

ica

l,

t=typ

ica

l)

Pelage

Totallength

Tail

Feet

Height

Craniallength

Odocoileusvirginianus

acapulcensis(Caton,

1877)

(aca)

A

capulco,

Guerrero,

M

xico

1394

195

385

677

224

Small,

frequently

reducedto

simplespikes

eveninadults

70t,90a

Varyingfrom

drabbishSn

uffBrowntoSayal

Brown,

thebandedhairs

tendingtoproducea

grizzledeffect


carminisGoldmanand

Kellogg,

1940(car)

B

otellasCan,

Sierra

d

elCarmen,northern

C

oahuila,

Mxico

1520

220

490

793

246

Moderately

spreadingwith

shortspikes

70?t,90?a

Adrabcoloredsubspecie

s.Upperpartsin

winterpelageamixtureo

fbrownishblackand

DrabGray.


couesi(CouesandYarrow,

1875)(cou)

R

anchoSantuario,

n

orthwesternDurango,

M

xico

1530

270

415

890

241

8-1

0spikes,with

thetendency

ofthemain

branchestoform

ovalinthefront

part

70t,90a

Brownishgrayordrabto

dullcinnamonor

cinnamonbuff.

Inwinter

pelagetheindividual

hairs,especiallyontheb

ack,areblackattips

withagrayishsub-termin

alband,

belowwhich

theyaredarkbrown


mexicanus(Gmelin,

1788)

(mex)

V

alleyofMxico,

Mxico

1550

235

410

915

241

4to8spikes,

branchescurve

tothefront

60?t,70?a

GeneralcolorationofupperpartsMummy

browntoCinnamonbrow

n


miquihuanensisGoldman

andKellogg,

1940(miq)

S

ierraMadreOriental,

n

earMiquihuana,

s

outhwesternTamaulipas,

M

xico

1530

270

420

820

247

similarto

texanobut

smallandshort

spikes

80?t,90?a

Darker,especiallyoverm

ediandorsalarea.

A

grizzledDrabsubspecies

;withconspicuous

brownishblackuppersid

eoftail;upperparts

inwornwinterpelageamixtureofSnuff

BrownandBuff.

Summerpelagemore

brownish.


nelsoniMerriam,

1898(nel)

S

anCristobal,highlands

o

fChiapas,Mxico

1346

170

371

658

224

Antlerswith

mainbeam

straighter,

directed

backward,

insteadof

curvingforward.

40?t,50?a

Adarkbrownishgrayhig

hmountain

subspecies

Appen

dix1.

Descriptionofthesubsp

eciesofwhite-taileddeerpresentinMexico,

basedonKellogg(1956)andVillarreal

(2009).Inparenthesisisshowt

heab

breviationsofsubspeciesusedinthefollowingtablesandgures.


26/26



oaxacensisGoldmanand

Kellogg,

1940(oax)

M

ountains15mileswest

o

fOaxaca,

Mxico

1340

170

362

750

230

4to8spikes,

branchescurve

tothefront

60?t,70?a

Presentagrizzledpattern

ofcoloration


sinaloaeJ.A.

Allen,

1903

(sin)

E

scuinapa,southern

S

inaloa,

Mxico

1490

223

415

820

234

Antlerslonger

withmore

numerouspoints

unknown

Varyingincolorfrom

nea

rSayalBrownto

drabbishdiluteSnuffBrown,mixedwithblack

overback


texanus(Mearns,1898)(tex)

F

ortClark,

Kinney

C

ounty,

Texas,USA

1829

254

420

1048

288

Thebiggest

sizedofantlers

ofallMexican

subspecies

100t,125a

Pelageshort,colorofupp

erpartsinwinter

pelageincliningtowardg

ray,usuallywith

grizzledpatternapproach

inggrayishlight

cinnamondrab


thomasiMerriam,

1898

(tho)

H

uehuetan,

Chiapas,

M

xico

1544

180

425

800

238

4to8spikes,

verticalbranches

slightlytoback

40?t,50?a

PredominanttoneofupperpartsTawnyor

CinnamonBrown


toltecus(Saussure,

1860)

(tol)

O

rizaba,

Veracruz,

M

xico

1440

71

4to8spikes,

verticalbranches

slightlytoback

40?t,50?a

Characterized

bydark

Odocoileusvirginianustruei

Merriam

1898(tru)

S

ibube,

Panam

1422

178

360

825

253

Antlersinadults

varyingfrom

simplespikesto

thosehavingtwo

orthreepoints

nearthetip.

40?t,50?a


veraecrucisGoldmanand

Kellogg,

1940(ver)

C

hijol,northernVeracruz,

M

xico

1560

220

400

800

247

smallandcloser

60?t,70?a

withtailusuallymoreorlessdistinctly

blackishbonebrownonuppersidesub

terminally.


yucatanensis(Hays,1872)

(yuc)

T

hroughoutYucatn

a

ndthesouthernpartof

M

xico

1470

228

365

647

251

Relatively

straight,simply

branched.

40?t,50?a

Palerless.

Finelygrizzled

woodbrownto

Mikadobrownupperparts

ontinuacinAppendix1...

Comparison of geographic distribution models of white-tailed deer Odocoileus virginianus...

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